Mortality of Apatania fimbriata (Insecta: Trichoptera) during embryonic, larval and adult life stages

1. Egg masses, oviposition site preferences, and abiotic and biotic factors causing mortality during embryonic, larval and adult life stages of Apatania fimbriata were studied. Laboratory investigations provided information on the temperature dependence of embryonic development, measured as an increase in egg volume.
2. A. fimbriata laid hemispherical egg masses, consisting of a transparent matrix containing a mean of 208 eggs. Egg masses were laid on stones situated just above the water surface in dark cavities in the stream bank.
3. Two hundred egg masses were mapped and individually monitored during embryonic development. There was no significant correlation between mortality during embryonic development and any of the abiotic parameters measured. First-instar larvae of Osmylus fulvicephalus consumed developing eggs, and chironomids preyed on newly hatched larvae.
4. A mean of seventy-two females emerged per metre of stream. Mortality during the 1993/94 life cycle was measured as a percentage of the potential number of eggs laid. Female mortality between emergence and oviposition was ≈ 80%. Eight per cent of individuals were lost during embryonic development. Larval mortality to emergence in 1994 was 11.3%. This indicates that the terrestrial life stage is probably decisive in the regulation of A. fimbriata populations.
5. Duration of embryogenesis at constant temperatures (4–20 °C) in the laboratory was described best by a negative exponential function. This species is cold stenothermal and there was no hatching success at 20 °C.
6. Egg volumes during embryonic development increased sigmoidally over time.     

Phylogeny of caddisflies (Insecta, Trichoptera)

Trichoptera are holometabolous insects with aquatic larvae that, together with the Lepidoptera, comprise the Amphiesmenoptera. Previous phylogenetic hypotheses and progress on our ongoing data collection are summarized. Fragments of the large and small subunit nuclear ribosomal RNAs (D1, D3, V4–5), the nuclear elongation factor 1 alpha gene and a fragment of mitochondrial cytochrome oxidase 1 (COI) were sequenced, and molecular data were combined with previously published morphological data. Equally and differentially weighted parsimony analyses were conducted in order to present a phylogeny of Trichoptera, including 43 of 45 families. Our phylogeny closely resembles that proposed by Herbert Ross with respect to the relationships among suborders, with a monophyletic Annulipalpia at the base of the tree, and a clade consisting of Spicipalpia plus a monophyletic Integripalpia. The monophyly of Spicipalpia is weakly supported in the combined equally weighted analysis, and Spicipalpia is paraphyletic in the differentially weighted analysis. Within Integripalpia, our phylogeny recovered monophyletic Plenitentoria, Brevitentoria and Sericostomatoidea. Leptoceroidea was unresolved in the equally weighted analysis and monophyletic in the differentially weighted analysis. Within Annulipalpia, we recovered a basal but paraphyletic Philopotamoidea and a monophyletic Hydropsychoidea.     

Mortality of the terrestrial adult and aquatic nymphal life stages of Baetis vernus and Baetis rhodani in the Breitenbach, Germany (Insecta: Ephemeroptera)

1. The mortality of Baetis vernus Curtis and Baetis rhodani Pictet during the terrestrial-aerial and aquatic life stages was studied at the Breitenbach near Schlitz, Hesse, Germany. The number of females emerging from the stream was recorded with emergence traps. To estimate mortality of females of both species during terrestrial life, numbers of emerging females were compared with numbers of females returning to the stream for oviposition, as shown by numbers of egg masses found in the stream. 2. Mortality of female B. vernus during their terrestrial life stage was 98.8%. It was 91.2 and 96.6%, respectively, during the first and second generations of B. rhodani. 3. To estimate the mortality of both sexes of B. rhodani during the aquatic life stage, the number of eggs laid by the first generation was compared with the number of adults emerging during the second generation. 4. Mortality during the aquatic phase (both sexes combined) of B. rhodani was 91.1%. 5. Mortality during the terrestrial life stages was at least as high, if not higher than during the aquatic stages. Evidently, there is a considerable export of organic material into the terrestrial environment around the stream. Mortality during terrestrial life may be an important regulator of population size.     

Differences in drifting of larval stages of Rhyacophila nubila (Trichoptera)

Larval drift of the trichopteran Rhyacophila nubila (Zett.) was studied in a West Norwegian river. The frequency distribution of different instars in the drift samples was significantly different compared with the benthic samples. The drift, measured as the number which passed a transect of the river, rised markedly in periods of high water discharge. Different instars can be arranged according to the extent in which they are drifting: 2>3>4>1>5. Instar 1 and 2 shared a neutral phototaxis. From the 2nd instar and on, the larvae became increasingly night-active with a culmination in the last instar which was highly night active, also confined by laboratory studies. It is suggested that changes in activity patterns is associated with a shift towards a more carnivorous way of life.     

Phylogeny of the Ecnomidae (Insecta: Trichoptera)

Ecnomidae are a family of seven previously accepted extant genera having a typical Gondwanan distribution, except one genus ( Ecnomus ) being widely distributed also in the Oriental and Palearctic regions. We analysed a molecular data set of 3379 characters representing the sum of four different protein-coding genes (COI, CAD, EF-1a and POL-II). Six equally most parsimonious trees were generated from the combined data set, distributed into two distinct islands. In all maximum parsimony (MP) trees the Ecnomidae is monophyletic when the genus Zelandoptila (Psychomyiidae) is included. The sister group to Ecnomidae including Zelandoptila is Pseudoneureclipsis , previously classified in the other families. This sister-group relationship contradicts earlier findings that the Polycentropodidae are the sister group to Ecnomidae. A Bayesian analysis resulted in a monophyletic Ecnomidae when accepting inclusion of Pseudoneureclipsis , which contradicts the results from the MP analysis by leaving Zelandoptila as the sister group to Ecnomidae including Pseudoneureclipsis . In the majority rule tree from this analysis Polycentropodidae form the sister group to Ecnomidae. We were not able to obtain a monophyletic Ecnomus due to the inclusion of Psychomyillodes . We conclude that the genus Zelandoptila or Pseudoneureclipsis probably belongs to the Ecnomidae, and that Psychomyiellodes and Ecnomus are synonyms. Three additional, as yet undescribed monotypic genera from Australia and New Caledonia remain to be erected in Ecnomidae.
© The Willi Hennig Society 2009.     

Effects of larval energetic resources on life history and adult allocation patterns in a caddisfly (Trichoptera: Phryganeidae)

Abstract 1. How populations respond to environmental change depends, in part, on the connection between environmental variance during early life stages and its effect on subsequent life‐history traits. For example, environmental variation during the larval stage can influence the life histories of organisms with complex life cycles by altering the amount of time spent in each stage of the life cycle as well as by altering allocation to life‐history traits during metamorphosis. 2. The effects of larval energetic resources on developmental timing, adult mass, fecundity, mating success, and allocation to adult body structures (thorax, abdomen, wings) were examined in an aquatic caddisfly (Agrypnia deflata Milne, Trichoptera: Phryganeidae). Larval energetic reserves were manipulated by removing larval cases just prior to pupation. In the first experiment, cases of all individuals were removed just prior to pupation; experimental individuals were required to build a new case whereas control individuals were allowed to re‐enter their case without building. In the second experiment, energy differences were maximised between the two treatments by supplementing the larval diet of the control group and removing cases and not supplementing the diet of the experimental group. 3. Male and female development time, adult mass, and female fecundity were not influenced by case removal or diet supplementation. In contrast, allocation to adult body parts indicated a trade‐off between abdominal and thoracic mass among case‐removal females, suggesting that, under larval resource stress, females adjust resource allocation during metamorphosis to alleviate potential negative impacts on clutch size. In addition, latency to copulation increased when cases were removed, indicating larval resource stress could influence male mating success. 4. This study suggests that, under larval energetic stress, the negative impacts on female reproduction might be mitigated by re‐allocating resources during metamorphosis, whereas male allocation strategies might not be as flexible as female strategies.     

Biological diversity of the Minnesota caddisflies (Insecta, Trichoptera)

The caddisfly fauna of Minnesota contains at least 277 species within 21 families and 75 genera. These species are based on examination of 312,884 specimens from 2,166 collections of 937 Minnesota aquatic habitats from 1890 to 2007. Included in these totals is my own quantitative sampling of 4 representative habitat types: small streams, medium rivers, large rivers, and lakes, from each of the 58 major Minnesota watersheds from June through September during 1999-2001. All species are illustrated herein, and their known Minnesota abundances, distributions, adult flight periodicities, and habitat affinities presented. Four species: Lepidostoma griseum (Lepidostomatidae), Psilotreta indecisa (Odontoceridae), and Phryganea sayi and Ptilostomis angustipennis (Phryganeidae) are added to the known fauna. An additional 31 dubious species records are removed for various reasons. Of the 5 determined caddisfly regions of the state, species richness per watershed was highest in the Lake Superior and Northern Regions, intermediate in the Southeastern, and lowest in the Northwestern and Southern. Of the 48 individual collections that yielded >40 species, all but 1 were from the Northern Region. Many species, especially within the families Limnephilidae and Phryganeidae, have appeared to decrease in distribution and abundance during the past 75 years, particularly those once common within the Northwestern and Southern Regions. Many species now appear regionally extirpated, and a few have disappeared from the entire state. The loss of species in the Northwestern and Southern Regions, and probably elsewhere, is almost certainly related to the conversion of many habitats to large-scale agriculture during the mid-20th century.     

Free amino acid concentrations in plasma of larval and adult hydropsychidae (Trichoptera)

Free amino acid concentrations in plasma of larval and adult Hydropsyche betteni and Cheumatopsyche pettiti were determined. Free amino acids in whole-body homogenates of these two species were also investigated.Larval haemoplasm of both species contained high concentrations of serine, proline, alanine, lysine, valine, threonine and arginine. Total free amino acid concentrations were consistently higher in C. pettiti larvae (33.1 mM) than in H. betteni larvae (24.6 mM).Haemoplasm from adult specimens of H. betteni contained high concentrations of serine and proline, followed by alanine, valine, lysine and histidine. Serine was found in very high concentrations in plasma from C. pettiti adults, followed by proline, glytamic acid, histidine and arginine. The mean total concentration of free amino acids was higher in C. pettiti adults (18.6 mM) than in H. betteni adults (13.3 mM).Comparison of free amino acids from haemoplasm and whole-body homogenates suggests that concentrations of free amino acids in body tissues differ from those present in plasma for these two insects.     

The genus Rhyacophila Pictet, 1834 (Insecta: Trichoptera) in Kosovo

The Balkan Peninsula is known for its high species diversity, but it is also one of the least known areas of Europe. Many previously inaccessible regions lack reliable faunistic records. There are especially few data available for the highly diverse aquatic insects of the region. To help fill the gap, we conducted an extensive survey to map the Trichoptera fauna of the Republic of Kosovo, an area where faunistic records are extremely scarce. Adult specimens were collected during 2009 and 2010, with light traps and entomologic nets. Of 84 Trichoptera species identified from the 21 investigated stations, 15 belong to the genus Rhyacophila. This makes the genus the most species-rich in Kosovo. Rhyacophila fasciata and R. tristis are widespread in many streams and rivers all over the country, while R. balcanica, R. trescavicensis, R. obtusa and R. bosnica have limited distribution. Ten Rhyacophila species presented in this paper are recorded for the first time on the area of Kosovo. We also provide new data for the disjunctive distribution of R. palmeni in south-eastern Europe, from Kosovo and Croatia.     

Global diversity of caddisflies (Trichoptera: Insecta) in freshwater

The not yet uploaded Trichoptera World Checklist (TWC) [], as at July 2006, recorded 12,627 species, 610 genera and 46 families of extant and in addition 488 species, 78 genera and 7 families of fossil Trichoptera. An analysis of the 2001 TWC list of present-day Trichoptera diversity at species, generic/subgeneric and family level along the selected Afrotropical, Neotropical, Australian, Oriental, Nearctic and Palaearctic (as a unit or assessed as Eastern and Western) regions reveals uneven distribution patterns. The Oriental and Neotropical are the two most species diverse with 47–77% of the species in widespread genera being recorded in these two regions. Five Trichoptera families comprise 55% of the world’s species and 19 families contain fewer than 30 species per family. Ten out of 620 genera contain 29% of the world’s known species. Considerable underestimates of Trichoptera diversity for certain regions are recognised. Historical processes in Trichoptera evolution dating back to the middle and late Triassic reveal that the major phylogenetic differentiation in Trichoptera had occurred during the Jurrasic and early Cretaceous. The breakup of Gondwana in the Cretaceous led to further isolation and diversification of Trichoptera. High species endemism is noted to be in tropical or mountainous regions correlated with humid or high rainfall conditions. Repetitive patterns of shared taxa between biogeographical regions suggest possible centres of origin, vicariant events or distribution routes. Related taxa associations between different regions suggest that an alternative biogeographical map reflecting Trichoptera distribution patterns different from the Wallace (The Geographical Distribution of Animals: With a Study of the Relations of Living and Extinct Faunas as Elucidating the Past Changes of the Earth’s Surface, Vol. 1, 503 pp., Vol. 2, 607 pp., Macmillan, London, 1876) proposed biogeography patterns should be considered. Anthropogenic development threatens biodiversity and the value of Trichoptera as important functional components of aquatic ecosystems, indicator species of deteriorating conditions and custodians of environmental protection are realised.     

Asymmetric larval head and mandibles of Hydrophilus acuminatus (Insecta: Coleoptera,Hydrophilidae): Fine structure and embryonic development

The larvae of a water scavenger beetle, Hydrophilus acuminatus, have strongly asymmetric mandibles; the right one is long and slender, whereas the left one is short and stout. The fine structure and embryonic development of the head capsule and mandibles of this species were examined using light and scanning electron microscopy, and asymmetries in shape were detected in these structures applying an elliptic Fourier analysis. The larval mandibles are asymmetric in the following aspects: whole length, the number, structure and arrangement of retinacula (inner teeth), and size and shape of both the molar and incisor regions. The larval head is also asymmetric; the left half of the head capsule is larger than the right, and the left adductor muscle of the mandible is much thicker than the right. The origin and developmental process of asymmetric mandibles were traced in developing embryos whose developmental period is about 270 h and divided into 10 stages. Mandibular asymmetries are produced by the cumulative effects of six stepwise modifications that occur from about 36% of the total developmental time onward. The significance of these modifications was discussed with respect to the functional advantages of asymmetries and the phylogeny of members of the Hydrophilidae.     

A multigene phylogeny and timeline for Trichoptera (Insecta)

The Trichoptera, or caddisflies, are traditionally split into two taxonomic subdivisions: the ‘retreat-making’ Annulipalpia and the ‘case-making’ Integripalpia (sensu Ross). The monophyly of these groups is well documented; however, the establishment of a third subdivision, ‘Spicipalpia’, and the positions of the five ‘spicipalpian’ families is much debated. In contrast to previous molecular studies using nuclear ribosomal RNA, a recent trichopteran study (using nuclear protein-coding genes) placed one of these ‘spicipalpian’ families, the free-living predatory Rhyacophilidae, as the sister taxon to the rest of Trichoptera, a result that has significant implications for both the understanding of trichopteran evolution and its timing. This paper sets out to investigate the relationships of Trichoptera using several newly sequenced genes, together with previously published gene sequences. This dataset is the largest trichopteran dataset to date, covering six independent genes and > 10 000 nucleotides, and containing 185 species representing 49 families. With all data included, likelihood and Bayesian analyses support a monophyletic Annulipalpia and a monophyletic Integripalpia, which includes the ‘spicipalpians’ as a paraphyletic grade at the base of this clade. However, an analysis of the protein-coding data alone using similar analytical methods recovers Rhyacophilidae as the most basal taxon in Trichoptera, with low support. A reanalysis correcting for nucleotide composition bias provides support for the placement of the ‘spicipalpian’ taxa as sister to the Integripalpia, consistent with the total data analysis, suggesting that the basal position of Rhyacophilidae in the uncorrected analysis could be (or is probably) an artefact of base composition. We find it likely that ancestral trichopterans made incipient cases and retreats, and these had independent origins as precocious pupal chambers. Molecular dating analysis in beast , using the birth-death model of speciation, with a relaxed-clock model of sequence evolution informed by 37 fossil constraints, suggests that the most recent common ancestor of Trichoptera appeared in the Permian (c. 275 Ma) in line with the first appearance of Trichoptera in the fossil record, and that vicariance explains the distribution of most trichopteran taxa. A new infraordinal name, Phryganides , is introduced for the tube-case-making families of Integripalpia.     

The evolutionary history of Trichoptera (Insecta): A case of successful adaptation to life in freshwater

The insect order Trichoptera (caddisflies) forms the second most species‐rich monophyletic group of animals in freshwater. So far, several attempts have been made to elucidate its evolutionary history with both morphological and molecular data. However, none have attempted to analyse the time frame for its diversification. The order is divided into three suborders – Annulipalpia, Integripalpia and ‘Spicipalpia’. Historically, the most problematic taxon to place within the order is ‘Spicipalpia’, whose larvae do not build traditional cases or filtering nets like the majority of the caddisflies. They have previously been proposed to be the sister group of all other Trichoptera or more advanced within the order, with equivocal monophyly and with different interordinal placements among various studies. In order to resolve the evolutionary history of the caddisflies as well as timing their diversification, we utilized fragments of three nuclear (carbamoylphosphate synthethase, isocitrate dehydrogenase and RNA polymerase II) and one mitochondrial (cytochrome oxidase I) protein coding genes, with 16 fossil trichopteran taxa used for time calibration. The ‘spicipalpian’ families are recovered as ancestral to all other caddisflies, though paraphyletic. We recover stable relationships among most families and superfamilies, resolving many previously unrecognized phylogenetic affinities amongst extant families. The origin of Trichoptera is estimated to be around 234 Ma, i.e. Middle – Late Triassic.     

The Micro-caddisflies of Lord Howe Island (Hydroptilidae: Trichoptera: Insecta)

Hydroptilid species are described from Lord Howe Island for the first time. All four species are in the endemic Australian genus Orphninotrichia, and all are endemic to Lord Howe Island. Modifications exhibited by these new species to the normally conservative Orphninotrichia features are suggestive of sympatric speciation and, for several, of a shift in larval niche.     

New long-horned caddisflies in Eocene Baltic amber (Insecta,Trichoptera)

We describe two new extinct caddisflies, Electroadicella kuenowi sp. nov. and Triaenodes simoni sp. nov., embedded in Baltic amber. Both species belong to the tribe Triaenodini and enlarge the family Leptoceridae to a remarkable number of 18 extinct Eocene species from Baltic amber. The extinct Electroadicella and the extant Triaenodes are possibly closely related, by a distinct synapomorphy, a recurved process from the basal plate of the inferior appendages of the ninth abdominal segment in the male genitalia.     

New records of polycentropodid caddisflies from Korea (Insecta: Trichoptera)

Four species of the family Polycentropodidae –Plectrocnemia baculifera, P. kusnezovi, Paranyctiophylax hjangsanchonus and Polyplectropus nocturnus– are recorded from Korea. Polyplectropus nocturnus is newly recorded from the Korean peninsula. The other three species are recorded from the Republic of Korea for the first time.     

Effects of lufenuron on Lobesia botrana (Lepidoptera: Tortricidae) egg, larval, and adult stages

The effect of the chitin synthesis inhibitor lufenuron was evaluated against different developmental stages of Lobesia botrana Den. & Schiff. (Lepidoptera: Tortricidae). Lufenuron fed to adults at 10 ppm reduced their fecundity and fertility, but it did not affect adult longevity. High activity was observed against L. botrana eggs with greater effect on 1-d-old eggs than on the other age classes and on eggs treated by direct contact rather than those laid on a previously treated surface. Eggs laid by treated adults showed the same effects during development as eggs treated by contact or those laid on a treated surface. Larvae that emerged from treated eggs could not perforate grape berries. Administered into the diet, lufenuron had a larvicidal effect, resulting in similar LC50 values for different instars: 0.07 ppm for first instars, 0.08 ppm for third instars, and 0.11 ppm for fifth instars. None of the larvae treated with sublethal concentrations throughout their life emerged as adults at the highest concentration (0.08 ppm), and only 70% emerged at the lowest concentration (0.0025 ppm).     

New Insects (Insecta: Trichoptera,Reculida, Eoblattida) from the Mesozoic of Asia

New caddisflies, reculid and eoblattid insects from the Mesozoic of Asia are described. Caddisflies of the families Philopotamidae (Mesoviatrix paradoxa gen. et sp. nov. and Kempia piotri gen. et sp. nov.) and Polycentropodidae (Polylongevus eskovi gen. et sp. nov.) from the Upper Jurassic–Lower Cretaceous of the Kempendyai locality in Yakutia are described. A short review and comparison of fossil members in these families are provided. New Gryllones insects, Shurabia tanga sp. nov., Sauk batkenicus gen. et sp. nov. from the Sauk Tan’ga locality (Lower Jurassic of Kyrgyzstan), Say kirgizicus gen. et sp. nov. from the Shurab III locality (Reculida: Geinitziidae; Lower or Middle Jurassic of Kyrgyzstan) Griphopteron iya sp. nov. (Eoblattida: Blattogryllidae) from Iya locality (Middle Jurassic of Russia) are described.     

New insects (Insecta: Trichoptera,Eoblattida) from the Lower Permian of Russia

The insect fauna of the Lower Permian Tyulkino locality in the Perm Region of Russia is considered. New members of the orders Trichoptera and Eoblattida are described. Members of the primitive family Microptysmatidae (suborder Protomeropina) are typical of the Lower and Middle Permian deposits of Eurasia and are represented by the two genera: Microptysmella Kukalová-Peck et Willmann, 1990 and Kamopanorpa Martynov, 1928. Kamopanorpa tyulkiana sp. nov. and K. tamarae sp. nov. (Trichoptera, Microptysmatidae) and Parakhosara kamica sp. nov. (Eoblattida, Megakhosaridae) are described.